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Downloaded from Bioscientifica.Com at 10/06/2021 02:24:44AM Via Free Access Superovulated with Twice Daily Injections of FSH (FSH-P: Table 1 Failure of sheep\p=n-\goathybrid conceptuses to develop to term in sheep\p=n-\goat chimaeras R. A. Gustafson , G. B. Anderson, R. H. BonDurant and G. R. Sasser department of Animal Science and department of Reproduction, University of California, Davis, CA 95616-8521 USA; and iDepartment of Animal Science, University of Idaho, Moscow, ID 83843, USA Six hybrid pregnancies were established: three in sheep\p=n-\goat chimaeras, one in a sheep\p=n-\ (sheep\p=n-\goat)hybrid chimaera and two in does. Pregnancies were monitored weekly by ultra- sonography and peripheral concentrations of pregnancy specific protein B (PSPB) were measured. Placental development as detected by ultrasonography appeared to be slower in hybrid-in-goat pregnancies than in hybrid-in-chimaera pregnancies, although this difference was not reflected in PSPB concentrations. Time of fetal death could not be predicted from PSPB concentrations. Chimaeras appeared to carry hybrid pregnancies longer than ewes and does usually carry hybrid pregnancies, but none was carried to term. Introduction measured (Allen, 1975). Pregnancy specific protein (PSPB) has been identified in maternal circulation during pregnancy in The domestic sheep, Ovis arles, and domestic goat, Capra hircus, ruminants, including cattle (Butler et al, 1982), sheep (Ruder et do not normally interbreed. Ewes usually will not conceive al, 1988), goats (Humblot et al, 1990) and deer (Wood et al, PSPB is localized in et and secreted when bred to a buck. Through the use of artificial insemination 1986). (Eckblad al, 1985) and embryo transfer, hybrid pregnancies have been established (Reimers el al., 1985) from the binucleate cells of the tropho- blastic ectoderm. The of PSPB has not in ewes, but they are lost by 4—6 weeks of gestation (Hancock biological function(s) et al, 1968; MacLaren et al, in press). Does will conceive when been determined but has been linked to immunosuppression et mated to a ram, but pregnancy is usually lost by 8 weeks of (Dunbar al, 1990). gestation (Warwick and Berry, 1949; Hancock et al, 1968; This study was undertaken to determine the outcome of Hancock and McGovern, 1968; McGovern, 1973a; MacLaren hybrid pregnancies in sheep-goat chimaeras, using secretion of et al, in press). Occasional sheep—goat hybrids born to does PSPB and ultrasonography to monitor pregnancy. have been reported (Bunch et al, 1976; Moore et al, 1981; Tucker et al, 1989). Dent et al. (1971) that loss of proposed Materials and Methods hybrid-in-goat pregnancies can be attributed to placental failure, possibly due to a maternal immune response (Hancock et al, 1968). Sheep-goat chimaeras provide a useful model to test this Animals theory because they are tolerant to both immunologically Establishment of hybrid pregnancies was attempted in six lines (MacLaren et 1992b; Gustafson et al, 1993). component al, female sheep—goat chimaeras produced in previous experiments If failure of is due to an immune hybrid-in-goat pregnancies (Roth et al, 1989; Ruffing, 1989), a sheep-(sheep-goat)hybrid the chimaera should be of carry¬ response, sheep—goat capable chimaera (Roth et al, 1989), two Finn ewes and two and but case. Alpine ing sheep, goat hybrid pregnancies, this is not the does. The chimaeras had previously resorbed caprine and female chimaeras were shown to lambs Eight sheep-goat carry carried to term ovine pregnancies (MacLaren et al, 1992a, b). to term, but were lost from day 23 to some goat pregnancies Hybrid embryo donors consisted of one multiparous and five time after day 40 (MacLaren et al, 1992a, b). Establishment of virgin Alpine does. Intraspecific pregnancies were obtained by hybrid in chimaeras was by pregnancies sheep-goat attempted mating three white-face ewes and an Alpine doe to a fertile ram Fehilly and Willadsen (1986); however, none developed to term and buck, respectively. and it is unclear whether pregnancy was ever established. If placental function is compromised in interspecific and hybrid pregnancies, production of placental products would be Synchronization, insemination and embryo transfer to be affected. Such an effect has been observed with expected All animals were synchronized by use of intravaginal mares mule where reduced con¬ carrying conceptuses greatly pessaries containing 40 mg fluorogestone acetate (Chrono-gest: centrations of chorionic are equine gonadotrophin (eCG) Intervet, Boxmeer) inserted for 10 days. Lutalyse (15 mg; 'Correspondence and reprint requests. Upjohn, Kalamazoo, MI) was administered at the time of Revised manuscript received 13 May 1993. pessary removal. Does used as hybrid embryo donors were Downloaded from Bioscientifica.com at 10/06/2021 02:24:44AM via free access superovulated with twice daily injections of FSH (FSH-P: Table 1. Results of hybrid embryo transfer to sheep—goat Shering, Kenilworth, NJ) in decreasing doses of 5.5, 4.4 and chimaeras and to ewes 3.3 mg, respectively, for 3 days beginning the day before removal. pessary Embryos Pregnancy Insemination of the donor does with 150-200 x 106 motile Animal transferred Outcome termination* ovine spermatozoa was performed by laparoscopy to one uterine horn near the bifurcation 50—54 h after pessary removal. 8702-SHC 1 < 125 For this procedure, does were sedated with diazepam (0.15 mg blastocyst Pregnant Day i.V.: Schein, Port NY) and the skin and 1 morula kg-1, Washington, 1 16-cell abdominal wall were anaesthetized by local infiltration of lido- S703-C 2 morulae Not pregnant caine Inc., NJ). Semen was collected (Elkins-Sinn, Cherry Hill, by 8704-C 1 morula Not pregnant and for 300 over a electroejaculation centrifuged 20 min at # 8806-C 2 blastocysts Pregnant < Day 89 discontinuous Percoli (Sigma, St Louis, MO) gradient consisting 8809-C 3 morulae Pregnant < Day 70 of 2 ml decreasing concentrations of 70%, 50% and 40% Percoli 8810-C 1 blastocyst delayed oestrusb solution diluted in 1 x basal medium (BM). The Percoli solution 1 morula consisted of 9:1 Percoll:10 x basal medium (10 x BM: 1.13 mol 8811-C 2 morulae Pregnant < Day 49 NaCl 1 0.03 mol KCI 1 0.2 mol Tris 1 0.1 mol 102-S 1 blastocyst Not pregnant " \ " \ " \ glucose 1, 103-S 2 morulae Not pregnant 0.05 mol sodium pyruvate 1~ , 0.1 mol CaCl2 I-, 500 iu penicillin G ml-1, 0.5 ng streptomycin sulfate ml-1). The sperm pellet was then washed twice with 1 x BM and the final suspen¬ aDay ultrasonography revealed a nonviable fetus or nonpregnant state. sion held at 37°C in 1 x BM supplemented with 0.5% BSA bDelayed return to oestrus, stood to be mounted by a ram on day 46. SHC: chimaera; C: chimaera; S: (Sigma) until use. sheep—hybrid sheep—goat sheep. Morula- to early blastocyst-stage hybrid embryos were col¬ lected at 5 after insemination to laparotomy days according serum to sheep PSPB was used with bovine PSPB (laboratory methods described by Anderson et al. (1981). Embryos were preparation R-37) as the labelled standard protein. The mean held in culture in Whitten's medium and (Whitten Biggers, sensitivity of the standard curve for two assays, expressed as 0.5% BSA 1% 1968) supplemented with and (v/v) antibiotic- two standard deviations from buffer control, was 156.2 pg per solution x , Grand under antimycotic (100 Gibco, Island, NY) assay tube (sample size was 200 pi per tube). Inter- and intra- St 5% dimethyl-polysiloxane (Sigma, Louis, MO) at 37°C in assay coefficients of variation were 8.05 and 5.92%, respect¬ Ovulation the C02 until transfer to recipients. in recipients ively. Serial dilutions (with virgin ewe serum) of pooled serum (chimaeras and ewes) was confirmed at laparotomy and one to of hybrid-in-goat and hybrid-in-chimaera pregnancies inhibited three were transferred to a uterine horn to a embryos ipsilateral binding of radiolabelled (I25I) bovine PSPB in a manner parallel luteum. does were established corpus Hybrid pregnancies in by to the standard curve. Serum samples from intraspecific ovine natural to a Barbados ram. mating and caprine pregnancies were previously tested (Willard et al, 1987) and also inhibited binding parallel to the standard curve. Pregnancy detection and monitoring Animals were observed daily for return to oestrus. Chimaeras Statistical analysis were housed with a vasectomized ram a wearing marking PSPB data were subjected to analysis of covariance using the harness from 15 onwards. Ewes and chimaeras that returned day SAS General Linear Models procedure (SAS, 1985). PSPB con¬ to oestrus were bred to rams to establish naturally intraspecific centrations (absolute values and increases above basal concen¬ and ovine Animals that did not show pregnancies, respectively. trations analysed separately) were used as the dependent of oestrus 26 of oestrus = 0) were signs by day (first day day variable and type of pregnancy (for example, sheep-in-sheep, examined real-time a 3.5 MHz trans- by ultrasonography using hybrid-in-chimaera) and day of pregnancy were independent abdominal twice first for detection probe weekly, pregnancy variables with day of pregnancy used as a covariance. PSPB and then until was lost or to within monitoring, pregnancy concentrations for the different types of pregnancy were com¬ 3—4 weeks of parturition. pared at day 24 using an unpaired t test. Day-24 PSBS con¬ Blood were collected from 18 to 32, samples daily days every centrations were compared because differences resulting from other from 34 to 60, twice from 62 to 89 day days weekly days different numbers of conceptuses are not detectable at this time and once weeks weekly from day 90 until 2-3 after parturition (Humblot et al, 1990). (or 1—2 weeks after pregnancy loss). Blood was collected by jugular venepuncture into 10 ml vacutainer tubes (Becton- Dickenson, Rutherford, NJ), allowed to coagulate at room Results temperature and centrifuged at 2500 g for 10 min.
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